Biogéographie de Madagascar, 1996 : 441-455

ORIGINS AND AFFINITIES OF THE SCORPION FAUNA OF MADAGASCAR

Wilson R. LOURENçO

Laboratoire de Zoologie (Arthropodes), M.N.H.N., 61 rue de Buffon 75005 Paris, FRANCE

ABSTRACT.- Since the appearence’of the classical publications of P o c o c ~ (1894) and KRAEPELIN (1905), the biogeography of the scorpions of Madagascar has been a complex puzzle: a very long period of separation between the great island and Africa was postulated by POCOCK (1894). Subsequent contributions (e.g., FAGE, 1929; MILLOT, 1948; PAULIEN, 1961; LEGENDRE, 1972), have increased considerably our knowledge of the scorpion fauna of Madagascar. However, many interpretations as to the origins and especially of the affinities of this fauna, have been impaired by lack of precise knowledge of the phylogeny of Madagascan scorpions. Mnities with African scorpions have been accepted by most authors for quite some time (VACHON, 1979). The apparent demonstration of aflFinities with the Indo-Malayan and Australian regions (FAGE, 1929; MLLOT, 1948; LEGENDRE, 1972), proved, however, to be the consequence of misinterpretation when a typical Gondwanian lineage was finally established by LOURENÇO (1983, 1985). This Gondwanian lineage was detected by the study of al1 the genera of Ischnuridae distributed between Australia and Cocos Island near the Central American Pacific Coast, suggesting that the present representatives of the Madagascar scorpion fauna are derived from protoelements of both the families Buthidae and Ischnuridae which were already present in Gondwanaland, previous to fragmentation and continental drift. Finally, recent study of new elements from Madagascar (LO~RENÇO, 1995, 1996a,b,c) reinforces the likelihood of affinities with Africa, and of new affinities with Sri Lanka and India.

KEY-W0RDS.- Scorpion, Madagascar, Archaic, Affinities, Continental drift, Biogeography

RESUME.-Depuis la publication des travaux classiques de P o c o c ~ (1894) et KRAEPELIN (1905), la biogéographie des Scorpions de Madagascar semble correspondre à un véritable puzzle: une très longue période de séparation et d’isolement entre la grande île et l’Afrique a été postulée par P o c o c ~ (1894). Des contributions subséquentes (e.g., FAGE, 1929; MILLOT, 1948; PAULIAN, 1961; LEGENDRE, 1972), ont considérablement augmenté les connaissances sur la faune des Scorpions malgaches. Néanmoins, plusieurs explications et/ou interprétations visant à élucider les origines et en particulier les affinites de cette faune, ont été biaisées par un manque de connaissance précise concernant la phylogénie des Scorpions malgaches. Des &nités avec la faune afiicaine ont été acceptées depuis fort longtemps par la plupart des auteurs (VACHON, 1979). L’apparente démonstration d’affinités avec les faunes des régions Indo-Malaise et Australienne (FAGE, 1929; MILLOT, 1948; LEGENDRE, 1972), se sont avérées être le résultat des mauvaises identifications, en particulier depuis qu’une lignée typiquement Gondwanienne a été établie par LOURENÇO (1983, 1985). Cette lignée Gondwanienne a pu être établie à partir de l’étude de tous les genres de la famille des Ischnuridae, répartis depuis l’Australie jusqu’à l’île de Cocos proche des côtes pacifiques de l’Amérique Centrale. Cette étude suggère que les représentants actuels de la faune scorpionique malgache dérivent des protoéléments des familles Buthidae et Ischnuridae, lesquels étaient déjà présents dans la Gondwanie avant la fragmentation et la dérive des masses continentales. Finalement, l’étude récente de plusieurs nouveaux éléments de la faune malgache (LOURENÇO, 1995, sous-presse), vient renforcer la véracité des affinités avec la faune africaine et démontre l’existence de nouvelles affinités avec le Sri Lanka et l’Inde.

In: W.R LOURENçO (éd.) Editions de I’ORSTOM, Paris

442 W.R. LOURENÇO

MOTS-CLES.- Scorpion, Madagascar, Archaique, Affinités, dérive continentale, Biogéographie

INTRODUCTION

Contributions to the knowledge of the scorpion fauna of Madagascar began with descriptions of species by GERVAIS (1844) POcoCK (1890, 1894), KRAEPELIN (1896, 1901) etc. FAGE (1929) produced the first monographic study of the group which was a comprehensive work for the period in which it was produced.

The peculiarities of the fauna of Madagascar have attracted the attention of several authors Who have attempted to reach biogeographical conclusions regarding their origins and affinities. However, since the classical publications of POCOCK (1894) and KRAEPELIN (1905), the biogeography of the scorpions of Madagascar has proved to be a complex puzzle. As long as 1894, POCOCK suggested that Madagascar had been isolated fi-om Africa for a very long period of time.

Subsequent contributors (FAGE, 1929; MILLOT, 1948; PAULIEN, 1961; LEGENDRE, 1972; VACHON, 1979), proposed interpretations of the origins and especially of the ailinities of Madagascar's fauna which have been biased by two major factors: (i) lack of precise knowledge regarding the phylogeny of Madagascan scorpions in relation to those of Africa and the Orient; and (ii) overestimation of the extent of contemporary knowledge of Madagascar's fauna. FAGE (1929) stated: (< since scorpion species are of large size and have been collected in al1 the regions of the island, it is possible to estimate that we know al1 the populations of the island as well as their relative density and distribution D. MLLLOT (1948) followed this up by claiming:

BIOGEOGWHY OF SCORPIONS 443

(ii) supporting new evidence by the introduction of new elements which reinforce the likehood of affinities with Afr-ica, and of new af€inities with Sri Lanka and India.

Table 1. Comparative composition of the present scorpion fauna of Madagascar and that of three different periods.

FAGE 1929

FAMILYBUTHIDAE

Grosphus Simon, 1880

Gr. madagascariensis (Gervais, 1844)

Gr. hirtus Kraepelin, 1901

Gr. Jmopiceus Kraepelin, 1901

Gr. bistriatus Kraepelin, 1901

Gr. limbatus Pocock, 1889

Gr. limbatus annulata Fage, 1929

Gr. grandidieri Kraepelin, 1901

Odonturus Karsch, 1879

O. baroni (Pocock, 1890)

Uroplectes Peters, 186 1

U. fischeri nigrocarinatus Kraepelin, 19 13

FAMILY SCORPIONIDAE Heteroscorpion Birula, 1903

H, opisthacanthoides (Kraepelin, 1895)

Opisthacanthus Peters, 1861

O. madagascariensis Kraepelin, 1894

Total: 2 families, 6 genera, 10 species

2 subspecies

MILLOT 1948

FAMILYBUTHIDAE

Gr. bistriatus

Gr. Javopiceus

Gr. hirtus

Gr. limbatus

Gr. limbatus annulata

0.baroni

Ufischeri nigrocarinatus

Babycurus Karsch, 1886

Babycurus graciiis Fage, 1946

Isometrus Hemprich & Ehrenberg, 1829

Isometrus maculatus @eGeer) - Mien

FAMILY SCORPIONIDAE H.opisthacanthoides

0.madagascariensis

Doubtful species (*)

Babycurus centrurimorphus Karsch, 1886

Isometrus madagassus Roewer, 1943

Total: 2 families, 7 genera, 10 species,

2 subspecies

(*) Grosphus madagascarienis

is not meutioned by Miilot.

VACHON 1969/79

FAMlLYBUTHIDAE

Gr. madagascariensis

Gr. hirtus

Gr. grandidieri

Gr. flavopiceus

Gr. limbatus

444 W.R. LOURENÇO

Gr. limbatus annulata

Gr. bistriatus

Gr. griveaudi Vachon, 1969

Tityobuthus baroni

Tityobuthus gracilis

FAMILY SCOWIONIDAE

H.opisthacanthoides

Omadagascariensis

' Doubtful species (*)

B.centrurimorphes

UJscheri nigrocarinatus

Total: 2 families, 4 genera and 12 species

(*) Lmadagassus is

not mentionned by Vachon

THIS STUDY

FAMLYBUTHIDAE

Gr. madagascariensis

Gr. hirtus

Gr. grandidieri

Gr. jlavopiceus

Gr. limbatus

Gr. annulata

Gr. bistriatus

Neogrosphus Lourenço, 1995

Neogrosphus griveaudi (Vachon)

Isometrus maculatus - Mien (*)

T. baroni

T.graci1i.v

Tityobuthus guillaumeti Lourenço, 1995

Tityobuthuspococki Lourenço. 1995

Tityobuthus Irrcileae Lourenço. 1996

hficrocharmus Lourenço, 1995 hficrocharmus

cloudsleythompsoni Lourenço, 1995

Microcharmus hauseri Lourenço, 1996

Pseudouroplectes Lourenço, 1995

Pseudouroplectes betschi Lourenço, 1995

FAMILY ISCHNURIDAE (**)

H. opisthacanthoides

O. madagascariensis

Opisthacanthus punctulatus Pocock, 1896

Paleocheloctonus Lourenço, 1996

Paleocheloctonus pauliani Lourenço, 1996

Total: 2 families, 9 genera, 19 species

(*) Isometrus madagussus is a synonym of

Lmaeulatus

(**) The genera of Scorpionidae were separated into two famifies Scorpionidae and Ischnuridae by LOURENÇO (1985).

BIOGEOGRAPHY OF SCORPIONS 445

1. POSSIBLES ORIGINS OF THE SCORPION FAUNA OF MADAGASCAR

As proposed in a recent paper (LOURENÇO, 1996d), a useful and didactic approach can be based on UDVARDY'S (1 98 1) division of biogeography into three spatio-temporal entities. In correlation with UDVARDY'S model, three major biogeographical events may tentatively be used to explain the distribution patterns currently observed. In the present paper, which provides an explanation of possible origins of the fauna of Madagascar, 1 shall limit myself to the first of the three scales: the phylogenetic or palaeobiogeographical scale. The other two scales are treated in a another paper (LOURENÇO, 1996d).

A 500myr-- ----------- 7

I PHYLOGENETIC SCALE: 1 PALBOBIOGEOGRAPHY 1

100 myr = I

200 myr -

I I

I 1 I

f I

10myr - i 2myr ---------

MILLENNIAL SCALE 1 I l m y r - PLEISTOCENE I I (POSTPLEISTOCENE) 1

BIOGEOGRAPHY I 1 I I

1 I I I

I 1000 yr -------

SECULARSCALE: 1 '

BIOGEOGRAPHY I I ECOLOGICAL l

I B I w 100 1000 10000 40000 km 'km km km

Fig. 1. Division of biogeography into the three spatio-temporal scales of Udvardy (modified after Udvardy, 198 1).

446 W.R. LOT-JRENÇO

The phylogenetic scale encompasses the evolutionary t h e of al1 biota and is limited in space only by the size of the earth (UDVARDY, 1981). On this scale only historical factors can be assumed since, for almost al1 .ecological conditions, data are largely or totally unknown. At this level the evolutionary process of biogeography is, to a considerable extent, a tributary of continental driR and plate tectonics. In relation to Madagascar the following biogeographical assumptions can be proposed: (i) the two major lineages of scorpions present today, i.e. the families Buthidae and Ischnuridae, are derived from pulmonate (Neoscorpionina) elements that originated in Laurasia and Gondwanaland during Pangean times; (ii) protobuthids were the dominant fauna during Pangean times, and the present distribution of Buthidae on al1 continental lands of the world is the result of a vicariant process resulting from the fragmentation of Laurasia and Gondwanaland; (iii) the protoischnurids certainly evolved in Gondwanaland during Pangean times, and the present distribution of the Ischnuridae clearly suggests a typical Gondwanian lineage (LOt.RENç0, 1985); (iv) a precise analysis of the present scorpion fauna of Madagascar reveals several primitive lineages which show only patchy distributions .in other continental lands such as Africa and India; and (v) the presence of these primitive lineages suggests a long period of isolation between Madagascar and other land masses, following the fragmentation of Gondwanaland and continental drift.

OCEAN

Fig. 2. Position of Pangea about 200 my B.P., and hypothetical ways of coastal colonisation by aquatic scorpions.

BIOGEOGRAPHY OF SCORPIONS

II. J?RESENT AFFINITIES OF THE SCORPION FAUNA OF MADAGASCAR

447

A. African affinities

Affinities with Afiican scorpions have been accepted by most authors for some time (FAGE, 1929; MILLOT, 1948; VACHON, 1979), and the present configuration of the fauna seems to indicate significant affinities with Afiica. The following examples will be discussed. (i) Grosphus, the Madagascan genus most rich in species, and Neogrosphus, a genus which probably evolved more recently fiom Grosphus, and which clearly show affinities with the Mican genera Odonturus and, in particular, with Uroplectes. These three genera share very remarkable sexual dimorphism of the pectines (FAGE, 1929). (ii) The recently discovered and described genus Pseudouroplectes also shows affinities with the genus Uroplectes. However, the morphology of the new genus suggests a more primitive lineage when compared, with both Uroplectes and Grosphus. These two genera evidently evolved later. Pseudouroplectes corresponds with primitive lineages, still present in Madagascar, which have vanished in other regions of the world. (iii) The genus Tityobuthus was poorly defined phylogeneticly until recently. The discovery of three new species, however, improves Our knowledge of this group. Many affinities with the Gondwanian genus Ananteris, present today in West Afiica and in South America are indicated. Ananteris seems to have had its centre of origin in the West part of Gondwanaland. This corresponds today with the South American continent. These two genera present several primitive characteristics e.g., small size and the absence of fulcra in the pectines (l). (iv) The genus Opisthacanthus is the most typical Gondwanian lineage observed today. Its present distribution ranges fiom Madagascar through Africa to South America with one population in the Island of Hispaniola in the Caribbean and another in the Island of Cocos in the Pacific. The centre of origin of this genus is probably in South Afiica (LOWNÇO, 1985). Previous statements concerning the presence of Opisthacanthus in the Indo-Malayan and Australian regions (FAGE, 1929; MILLOT, 1948; LEGENDRE, 1972) have proved to be the consequence of misinterpretation. As stated by FAGE (1929) and by MILLOT (1948), the species of Opisthacanthus in Madagascar are closely allied to Opisthacanthus davydovi Birula, of the Am Islands, Indonesia. Subsequent studies, including one of the type specimen of 0.datydovi by LOURENÇO (1983) have revealed that this species belongs in fact to the genus Liocheles (2), a common genus in the Indo-Malayan and Australian regions. This genus, however, is absent fiom Madagascar and Mica. Opisthacanthus also presents some affinities with the genera Cheloctonus fiom Mica and Chiromachetes fiom India.

The absence of fülcra is observed in at least one species of Tiwobuthus

Presumably Fage nor Millot never examined Birnla's type. Moreover, these authors seem totally to ignore the genus Liocheles (previously Hormurus) in their studies.

448 W.R. LOURENÇO

AfEnities with the latter, however, have, to be established better. (v) The recent discovery of the genus Paleocheloctonus in the southwest region of Madagascar (LOWNçO, 1996c), clearly reinforces the claim for affinities with African scorpions. This genus is closely associated with the South African genus Cheloctonus. It represents, however, a more primitive lineage, which probably survived in Madagascar after the separation with Mica.

Fig. 3. Gondwanian distribution (black circles ) of scorpions of the genus Opisthacanthus.

B. Oriental affinities

With the discovery of the new genus Microcharmus in the North-east region of Madagascar, clear &nity is established with the genus Charmus of India and Sri Lanka. Both Microcharmus and Charmus represent primitive lineages whose characteristics show that they are among the very first modern buthid scorpions. No other evidence is at present available to indicate affinities between Madagascar and the Tndo-Malayan region,

BIOGEOGRAPHY OF SCORPIONS 449

and no evidence is available to suggest affinities between Madagascar and the Australian region.

Fig. 4. Distribution of the genus Ananteris in tropical America (1) and Afkica (2), and of the related species of Tityobuthus in Madagascar (3).

C. Particular cases

The phylogenetic and biogeographical position of some of the scorpions of Madagascar and the islands of the Indian Ocean are a complex puzzle. (i) The genus Heteroscorpion is monotypic and endemic to the north of Madagascar; some affinities can be suggested between it and the South Afi-ican genus Hadogenes. However, many common characteristics can only be due to convergence. The genus Heteroscorpion certainly represents a very old lineage, much older than Hadogerzes. (ii) The genus Chiromachus, another. monotypic genus, is only known from the Seychelles and Mauritius. Its absence from Madagascar is dificult to explain. Some affinities can be seen between this genus and the genus Chzronzachetes of India.

450 W.R, LOURENçO

In conclusion, the main event which determined the original biogeographical pattern of the scorpion fauna of Madagascar, on the palaeogeographic scale, was the fragmentation of Gondwanaland and subsequent continental drift. Difficulties in explaining the patchy distribution of the genera Heteroscorpion and Chiromachzs point not only to the great geological age of most groups, but also to the relict biogeographical patterns which they exhibit today.

Fig. 5. Distribution of Charmus in India and Sri Lanka (black circles) and of Microcharmzts in Madagascar (black circle with star).

m. SIGNIFICANT GAPS IN THE SCORPION FAUNA OF MADAGASCAR

As pointed out by PAULIAN (1952), the fauna of Madagascar has not only to be seen in connection with its remarkable originality, but also in respect to the gaps it presents. If two of the major lineages represented in Afi-ica and in the Indo-Malayan region, i.e. the families Buthidae and Ischnuridae, are also present in Madagascar, others are not. Two major gaps can be observed in Madagascar: (i) the family Scorpionidae which is present in Africa and in the Indo-Malayan region, and (ii) the family Chaerilidae which occurs in the Indo-Malayan region.

BIOGEOGRAF'HY OF SCORPIONS 45 1

As long ago as 1894, POCOCK stated that the apparent absence from Madagascar of large Afrcan genera of Scorpionidae was signifrcant, and he concluded that this indicated that this family had made its way to Central and South Afi-ica when the separation of Madagascar and Africa had been concluded. In fact, the Scorpionidae represent a more recent lineage than do either the Buthidae or the Ischnuridae. They probably evolved in some part of what is today North Africa, and later colonised the South and East (via the Middle East), displacing the Ischnuridae to the South. This is implied by the biogeographical patterns observed today. By the time they had colonised most of Africa, the separation of Madagascar had already taken place so they were unable to reach it (see also HEWITT, 1923).

Fig. 6. Distribution of the genera Heteroscorpion (black triangle), Chiromachus (black squares), Chiromachetes (black.. stars) and Iomachus (black circles), in Madagascar, Seychelles, India and Afrca.

With regard to the family Chaerilidae, LAMORAL (1980) proposed a possible explanation of its present pattern of distribution. According to this author, the protoelements of the Chaeriloids evolved in Laurasia, and the restriction of the Chaerilidae to the Oriental faunal region suggests that they are a relict of an eastern

452 W.R. LOURENÇO

Laurasian element that moved in after the conjunction with India. For this reason it seems logical to assume that the family Chaerilidae doen not exist in Madagascar.

W . DIVERSITY AND ENDEMISM OF THE SCORPION FAUNA OF MADAGASCAR

Subsequent to the biogeographical approach developed in the previous sections, it may be usefùl consider the diversity and endemicity of the scorpion fauna of Madagascar. When compared with other well studied regions of the world. (Table 11) which have approximately the same surface area or are smaller, Madagascar appears to possess a poor scorpion fauna. The total number of native species (19) is remarkably low when compared with Baja California (61), or with countries having a much smaller surface area, such as Ecuador (36). The number of families is also low; but other regions such as Imeri in BraziliadVenezuelan Amazonia, or Imataca in Guayana, which are likewise typical centres of endemicity, also contain only a small number of families. The total number of genera in Madagascar, however, seems significantly greater (8) when compared with that of Ecuador (S), or even of Baja California (1 1). Moreover, the inventory of species in Baja California and Ecuador is more complete than it is in Madagascar. Therefore, it seems probable that many new species and possibly new genera will be discovered. and described from Madagascar in the fùture. In recent additions to the scorpion fauna of Madagascar (LOUREN~O, 1995, 1996a,b,c,d,), 7 species have been described or revalidated and 4 new genera were described. This suggests that one of the particularities of the scorpion fauna of Madagascar is the presence of many generical lineages which apparently are not very rich in species.

As far as endemicity is concerned, the picture is quite different. Madagascar appears to be a region of the world which possesses one of the highest (if not the highest) level of endemic species of scorpions (and of other taxa also; see PAULIAN, 1952). Of the 19 native species of scorpions (3) al1 are endemic to Madagascar. Seven genera out of 8 are also endemic to Madagascar which corresponds to 87.5%.

CONCLUSIONS

In conclusion, although this study is only preliminary, the following characteristics

1. The majority of taxa found in Madagascar correspond with primitive or archaic

2. Most genera appear to be poor in species. However, it is reasonable to expect the fùture discovery of several new species ,in some genera of micro-scorpions such as Tityobuthus, Microcharmus and Pseudouroplectes.

of the scorpion fauna of Madagascar can be suggested:

lineages which no longer exist in most other regions of the world.

Isometrus muculutus an imported species, only occasionaly found in some coastal areas is not included here

BIOGEOGRAPHY OF SCORPIONS 453

3. The number of recorded species is apparently of little importance, however, and

4. The total number of genera is significant, even when compared with other well- studied regions of the world.

5 . The most remarkable characteristic of the scorpion fauna of Madagascar is the impressive level of endemicity, both in species and in genera. This supports the hypothesis of the very early isolation of the island fiom other land masses.

the faunistic inventories which have been carried out so far are largely incomplete.

Table JI and Graph 1. Comparative values of diversity and endemism observed in the Madagascan scorpion community and in several other well-studied regions of

the world.

Families Madagascar 2

Guayana 3 lmeri ' 2

. , lmataca 2 Ecuador 4

Paraguay 2 B. California 5

Genera 9 8 6 6 8 6 11

Species Endemic spp. 19 19 34 23 14 13 18 15 36 24 12 2 61 46

1 O0 90

%endemie 100.0 76.5 93. O 83.3 66.7 17.0 75.4

0 Families Genera

H Species

0 Endemic spp. %endemic

9 m

454 W.R. I..OURENÇO

ACKNOWLEDGEMENTS

In this article, 1 benefited from the comments of Profs. M.D.F. Udvardy and W.D. Shepard, California State University; J.L. Cloudsley-Thompson, University College London; Ch.P. Blanc, Université Montpellier 3 .1 thank al1 of them.

REFERENCES

BLACKBURN, T.M. & K.J. GASTON: 1994. Animal body size distributions change as more species are described. Proc. R. Soc. London, B, 257: 293-297.

FAGE, L., 1929. Les Scorpions de -Madagascar. Faune des Colonies fiançaises 3. Soc. Edit. Géogr. Marit. Colon., Paris: 637-694.

FAGE, L., 1946. Complèment à la faune des Arachnides de Madagascar. Bull. Mus. natl. Hist. nat., ~ Paris, 2è sér., 18(3): 256-267.

FENCHEL, T., 1993. There are more small than large species? Oikos, 68(2): 375-378.

GERVAIS, P., 1844. Remarques sur la famille des Scorpions. Archs. Mus. Hist. nat., Paris, 4: 201-240.

HEWITT, J., 1923. Remarks on the distribution of animals in South Afiica. South African J. Sci., 20:

KRAEPELIN, K., 1896. Neue und Weniger bekannte Scorpione. Jahr. Hamb. Wiss. Anst., 13: 121-146. KRAEPELIN, K., 1901. Catalogue des Scorpiones des collections du Muséum d'Histoire Naturelle de

96-123.

Paris. Bull. Mus. Hist. nat. Paris, 7: 265-274.

W P E L I N , K., 1905. Die Geographische Verbreitung der Scorpione. Zool. Jahr. Abt. Syst. 22: 321- 364.

LAMORAL, B.H., 1980. A reappraisal of suprageneric classification of recent scorpions and of their zoogeography. Proc. VIII, Inter. Arach. Kongr., Wien, 1980: 439-444.

LEGENDRE, R., 1972. Les Arachnides de Madagascar. In: Biogeography and ecology of Madagascar, G. Richard-Vindard & R. Battistini (Eds). pp. 427-457. W. Junk, The Hague.

LOURENçO, W.R., 1983. Considérations sur les genres Liocheles, Ischnurus, Opisthacanthus, Hormurzts, Hadogenes et Chiromachus appartenant à la sous-famille des Ischnurinae (Scorpiones : Scorpionidae). Ann. Natal Mus., 25(2): 403-411.

LOURENçO, W.R., 1985. Essai d'interprétation de la distribution du genre Opisthacanthus (Arachnida, Scorpiones, Ischnuridae) dans les régions néotropicale et afkotropicale. Etude taxinomique, biogéographique, évolutive et écologique. Thèse de Doctorat d'Etat, Univ. Pierre et Marie Curie: 287p.

LOURENçO, W.R., 1995. Description de trois nouveaux genres et de quatre nouvelles espèces de Scorpions Buthidae de l'île de Madagascar. Bull. Mus. natn. Hist. nat., Paris, 4e sér., 17(1-2): 79-90.

LOURENÇO, W.R., 1996a. Microcharmus hauseri, nouvelle espèce de Scorpion de Madagascar (Scorpiones, Buthidae). Rev. suisse Zool., 103(2):

BIOGEOGRAF'HY OF SCORPIONS 455

LOURENçO, W.R., 1996b. Tityobuthus lucileae, nouvelle espèce de Scorpion de Madagascar (Scorpiones, Buthidae). Boll. Mus. Region. Sci. Nat., Torino.

LOURENçO, W.R., 1996c. Scorpions. In: Faune de Madagascar. M.N.H.N., Paris.

LOURENçO, W.R., 1996d. The biogeography of scorpions. Rev. suisse 2001. (hors sér.).

MILLOT, J., 1948. Revue générale des Arachnides de Madagascar. Mém. Inst. Sci. Madagascar, Sér. A l(2): In: Mém. Soc. Biogéogr. (1953): 127-145.

PAULIAN, R., 1952. Esquisse du peuplement entomologique de Madagascar. Mém. Inst. Sci. Madagascar, Sér. E l(1): In: Mém. Soc. Biogéogr. (1953): 337-358.

PAULIAN, R., 1961. La zoogéographie de Madagascar et des îles voisines. In: Faune de Madagascar, Inst. Rech. Sci., Tananarive, 485p.

POCOCK, R.I., 1890. A revision of the genera of scorpions of the family Buthidae, with descriptions of

POCOCK, R.I., 1894. Scorpions an their geographical distribution. Natural Science, 4(27): 353-364.

UDVARTIY, M.D.F., 1981. The riddle of dispersal: dispersal theories and how they affect vicariance biogeography. In Vicariance biogeography: a critique. G.Nelson & D.E.Rosen (Eds.) Columbia Univ. Press. pp. 6-39.

some South-African species. Proc.. Zool. Soc. London, 11: 114-141.

VACHON, M., 1969. Grosphus griveaudi, nouvelle espèce de Scorpion Buthidae Malgache. Bull. Mus. natn; Hïst. nat., Paris 2è sér. 4(2): 476-483.

VACHON, M., 1979. Remarques biogéographiques sur la faune des Scorpions de Madagascar à propos de l'utilisation de caractères trichobothriotaxiques permettant la distinction des genres Odonturus Karsch, 1879 et Tityobuthus Pocock, 1893. C.R.Vè. Coll. Arach., M: 217-224.